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The dwarf crocodile (''Osteolaemus tetraspis''), also known as the African dwarf crocodile, broad-snouted crocodile (a name more often used for the Asian
mugger crocodile The mugger crocodile (''Crocodylus palustris'') is a medium-sized broad- snouted crocodile, also known as mugger and marsh crocodile. It is native to freshwater habitats from southern Iran to the Indian subcontinent, where it inhabits marshes, ...
) or bony crocodile, is an African
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
that is also the smallest
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
(living) species of crocodile.


Taxonomy and etymology

The second species has had a somewhat convoluted
taxonomical In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
history. It was first described as ''Osteoblepharon osborni'' by
Schmidt Schmidt may refer to: * Schmidt (surname), including list of people with the surname * Schmidt (singer) (born 1990), German pop and jazz singer * Schmidt (lunar crater), a small lunar impact crater * Schmidt (Martian crater), a List of craters on ...
in 1919, based on a few specimens from the Upper
Congo River Basin The Congo River ( kg, Nzâdi Kôngo, french: Fleuve Congo, pt, Rio Congo), formerly also known as the Zaire River, is the second longest river in Africa, shorter only than the Nile, as well as the second largest river in the world by discharge ...
in what is now the
Democratic Republic of Congo The Democratic Republic of the Congo (french: République démocratique du Congo (RDC), colloquially "La RDC" ), informally Congo-Kinshasa, DR Congo, the DRC, the DROC, or the Congo, and formerly and also colloquially Zaire, is a country in ...
. However,
Inger Inger may refer to: People * Inger (given name), a list of people * Inger, the main character of Hans Christian Andersen's fairy tale ''The Girl Who Trod on a Loaf'' * Robert F. Inger (1920–2019), American herpetologist * Stella Inger, America ...
in a 1948 paper found the specimens wanting of characteristics that would justify a generic separation from ''Osteolaemus'' and referred the specimens to '' Osteolaemus osborni''. In 1961, it was reduced to subspecies rank. A study of
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
published in 2007, and studies of DNA in 2009, 2013 and 2015 indicate that three distinctly different populations of ''Osteolaemus'' may merit full species recognition. These are ''O. tetrapis'' (
Central Africa Central Africa is a subregion of the African continent comprising various countries according to different definitions. Angola, Burundi, the Central African Republic, Chad, the Democratic Republic of the Congo, the Republic of the Congo, ...
, except the Congo River Basin), ''O. osborni'' (Congo River Basin), and a third possibly unnamed species (
West Africa West Africa or Western Africa is the westernmost region of Africa. The United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, Mali, Maurit ...
). Uncertainty exists for the population in
Nigeria Nigeria ( ), , ig, Naìjíríyà, yo, Nàìjíríà, pcm, Naijá , ff, Naajeeriya, kcg, Naijeriya officially the Federal Republic of Nigeria, is a country in West Africa. It is situated between the Sahel to the north and the Gulf o ...
(between ''O. tetrapis'' and the possibly unnamed West African species) as it has not been studied. A fourth
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
was found in a study of captives in 2013, but where members of this clade live in the wild is unclear. In some regions the species may come into contact. For example,
Cameroon Cameroon (; french: Cameroun, ff, Kamerun), officially the Republic of Cameroon (french: République du Cameroun, links=no), is a country in west-central Africa. It is bordered by Nigeria to the west and north; Chad to the northeast; the C ...
is home to both ''O. tetrapis'' and ''O. osborni''.


Etymology

The generic name, ''Osteolaemus'', means "bony throat", and is derived from the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
όστεον (bone) and λαιμός (
throat In vertebrate anatomy, the throat is the front part of the neck, internally positioned in front of the vertebrae. It contains the pharynx and larynx. An important section of it is the epiglottis, separating the esophagus from the trachea (windpipe ...
). The genus was named as such due to the osteoderms found among the scales in the neck and belly. The
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
, ''tetraspis'', means "four shields", and derives from the Ancient Greek τετρα (four) and ασπίς (shield), as the back of the neck has four large, shield-like scales.


Phylogeny

A 2018
tip dating Tip dating is a technique used in molecular dating that allows the inference of time-calibrated phylogenetic trees. Its defining feature is that it uses the ages of the samples to provide time information for the analysis, in contrast with traditio ...
study by Lee & Yates simultaneously using morphological, molecular (
DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
), and
stratigraphic Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithostra ...
(
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
age) data established the inter-relationships within
Crocodylidae Crocodiles (family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to include all extant mem ...
. In 2021, Hekkala ''et al.'' were able to use
paleogenomics Paleogenomics is a field of science based on the reconstruction and analysis of genomic information in extinct species. Improved methods for the extraction of ancient DNA (aDNA) from museum artifacts, ice cores, archeological or paleontological site ...
, extracting DNA from the extinct ''
Voay ''Voay'' is an extinct genus of crocodile from Madagascar that lived during the Late Pleistocene to Holocene, containing only one species, ''V. robustus''. Numerous subfossils have been found, including complete skulls, noted for their distinctiv ...
'', to better establish the relationships within Crocodylidae, including the subfamilies
Crocodylinae Crocodylinae is a subfamily of true crocodiles within the family Crocodylidae, and is the sister taxon to Osteolaeminae ( dwarf crocodiles and slender-snouted crocodiles). Taxonomy Crocodylinae was cladistically defined by Christopher Brochu in ...
and
Osteolaeminae Osteolaeminae is a subfamily of true crocodiles within the family Crocodylidae containing the dwarf crocodiles and slender-snouted crocodiles, and is the sister taxon to Crocodylinae. Taxonomy Osteolaeminae was named by Christopher Brochu in ...
. The below
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
shows the results of the latest study:


Characteristics

Dwarf crocodiles attain a medium adult length of , though the maximum recorded length for this species is . Adult specimens typically weigh between , with the largest females weighing up to and the largest males weighing . This makes it the smallest living crocodile species, although the
Cuvier's dwarf caiman Cuvier's dwarf caiman (''Paleosuchus palpebrosus'') is a small crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago ...
(''Paleosuchus palpebrosus''), a member of the family
Alligatoridae The family Alligatoridae of crocodylians includes alligators, caimans and their extinct relatives. Phylogeny The superfamily Alligatoroidea includes all crocodilians (fossil and extant) that are more closely related to the American alligator ...
, is smaller at up to about . If the Congo dwarf crocodile (''O. osborni'') is recognized as a valid species, it would be both the smallest crocodile and the smallest
crocodilian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
since it does not surpass . Adults are all dark above and on their sides, while the underside is yellowish with black patches. Some individuals living in the caves of Abanda, Gabon, displayed orange patches, apparently due to alkaline bat
guano Guano (Spanish from qu, wanu) is the accumulated excrement of seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. G ...
that erodes the skin of the crocodile. Juveniles have a lighter brown banding on body and tails and yellow patterns on the head. As a result of its small size and heightened vulnerability to predation, this species of crocodile has a heavily armoured neck, back, and tail and also has
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct ...
on its belly and underside of neck. ''Osteolaemus'' has a blunt short snout, as long as it is wide, similar to that of a Cuvier's dwarf caiman, probably a result of occupying a similar
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
. The
dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
consists of four
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
ry teeth, 12 to 13 on the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
, and 14 to 15 on the
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
bone. ''O. t. tetraspis'' has lighter colours, a more pointed, upturned snout, and more body armour than ''O. t. osborni''.


Distribution and habitat

Dwarf crocodiles range across
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
regions of
Sub-Saharan Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the African co ...
West Africa and Central Africa. Such a distribution greatly overlaps with that of the
slender-snouted crocodile ''Mecistops'' is a genus of crocodiles, the slender-snouted crocodiles, native to sub-Saharan Africa. Taxonomy and etymology Traditionally placed in ''Crocodylus'', recent studies in DNA and morphology have shown that it is in fact basal to '' ...
, encompassing countries as far west as
Senegal Senegal,; Wolof: ''Senegaal''; Pulaar: 𞤅𞤫𞤲𞤫𞤺𞤢𞥄𞤤𞤭 (Senegaali); Arabic: السنغال ''As-Sinighal'') officially the Republic of Senegal,; Wolof: ''Réewum Senegaal''; Pulaar : 𞤈𞤫𞤲𞤣𞤢𞥄𞤲𞤣𞤭 ...
, reaching
Uganda }), is a landlocked country in East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territor ...
in the east, and ranging as southerly as
Angola , national_anthem = " Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordina ...
. The last confirmed record from Uganda was in the 1940s, but whether the species, which is easily overlooked, still survives there is unclear (it was always marginal in this country, only occurring in the far southwest). Dwarf crocodiles live from lowlands to mid-altitude in streams, small rivers, swamps, pools and
mangrove A mangrove is a shrub or tree that grows in coastal saline water, saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evoluti ...
, but generally avoid main sections of large rivers. Most of their range is within forested regions, but it may extend into more open regions where the streams or river are well-shaded. Unlike most crocodiles, dwarf crocodiles only rarely bask in the sun. During the night they may move some distance from water on land. Reports exist of dwarf crocodiles in isolated pools in the
savannah A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the Canopy (forest), canopy does not close. The open canopy allows sufficient light to rea ...
. Dwarf crocodiles living long-term in caves are known from western
Gabon Gabon (; ; snq, Ngabu), officially the Gabonese Republic (french: République gabonaise), is a country on the west coast of Central Africa. Located on the equator, it is bordered by Equatorial Guinea to the northwest, Cameroon to the north ...
, which stand out as an isolated genetic group.


Biology and behaviour

The dwarf crocodile is a timid and mainly
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
reptile that spends the day hidden in pools or burrows, although it occasionally may be active during the day. Foraging is mainly done in or near the water, although it is considered to be one of the most
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
species of crocodilian and may expand the feeding pattern to land in extensive forays, especially after rains. Dwarf crocodiles are
generalist A generalist is a person with a wide array of knowledge on a variety of subjects, useful or not. It may also refer to: Occupations * a physician who provides general health care, as opposed to a medical specialist; see also: ** General pract ...
predators and have been recorded feeding on a wide range of small animals such as fish, crabs, frogs,
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s, insects, lizards, water birds, bats and
shrew Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to different fa ...
s. In a study in the Democratic Republic of the Congo the primary food item was fish, and in a study in Nigeria the primary food items were gastropods and crabs. In the Congo there is a level of seasonality in its diet, changing from fish in the wet season to crustaceans in the dry season, when fish are less available. Plant material has also been found in the stomach of dwarf crocodiles, but it is suspected that this is ingested by accident. They can survive for relatively long periods without food. During the dry season, dwarf crocodiles often retreat to deep holes. True to its solitary, nocturnal nature, a dwarf crocodile digs out a burrow in which to hide and rest during the day, which can sometimes have a submerged entrance. An individual lacking the right conditions to do so usually lives between tree roots that hang over the ponds where it lives.


Reproduction

Interacting closely only in
breeding season Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
, female dwarf crocodiles build their nest mounds at the beginning of the wet season, which spans May and June. The nest, situated near the water, is a mound of wet, decaying vegetation that incubates the
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
s due to the heat generated by the decomposition of the plant material. A small number of eggs is laid, usually about 10, though in extreme cases up to 20, and they incubate in 85 to 105 days. Hatchlings measure 28 cm when emerging from the eggs. The female guards the nest during the incubation period, and after the eggs hatch, she watches over the young for an unknown period of time, as young can be eaten by a great range of predators (birds, fish, mammals and reptiles, including other crocodiles).


Conservation

The dwarf crocodile is considered vulnerable by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
, and it is listed on Appendix I of
CITES CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...
. It is a little-known species, so unlike their more studied relatives, conservationists are often not as aware of how their populations are faring under the growing human pressure over the
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
s where they abide. Survey data, when available, show some degree of decline, either by hunting for
bushmeat Bushmeat is meat from wildlife species that are hunted for human consumption, most often referring to the meat of game in Africa. Bushmeat represents a primary source of animal protein and a cash-earning commodity for inhabitants of humid tropi ...
or habitat loss due to
deforestation Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated d ...
. However, it is a widely spread, and presumably numerous overall. In some regions the populations remain healthy, but in others (such as
Gambia The Gambia,, ff, Gammbi, ar, غامبيا officially the Republic of The Gambia, is a country in West Africa. It is the smallest country within mainland AfricaHoare, Ben. (2002) ''The Kingfisher A-Z Encyclopedia'', Kingfisher Publicatio ...
and
Liberia Liberia (), officially the Republic of Liberia, is a country on the West African coast. It is bordered by Sierra Leone to Liberia–Sierra Leone border, its northwest, Guinea to its north, Ivory Coast to its east, and the Atlantic Ocean ...
) it has seriously declined and may risk
extirpation Local extinction, also known as extirpation, refers to a species (or other taxon) of plant or animal that ceases to exist in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinct ...
. Dwarf crocodiles occur in several protected reserves. Though some skins are used in local manufacturing of
leather Leather is a strong, flexible and durable material obtained from the tanning, or chemical treatment, of animal skins and hides to prevent decay. The most common leathers come from cattle, sheep, goats, equine animals, buffalo, pigs and hogs, ...
products, they are of poor quality, so little interest is shown in
captive breeding Captive breeding, also known as captive propagation, is the process of plants or animals in controlled environments, such as wildlife reserves, zoos, botanic gardens, and other conservation facilities. It is sometimes employed to help species that ...
or a sustainable use program. In contrast, they are sometimes hunted for food and part of the bushmeat trade. Dwarf crocodiles are widely kept and bred in zoos. Based on a study of individuals kept in
AZA Aza or AZA may refer to: Places *Aza, Azerbaijan, a village and municipality *Azadkənd, Nakhchivan or Lower Aza, Azerbaijan *Aza, medieval name of Haza, Province of Burgos, Spain *Aźa, a Tibetan name for the Tuyuhun kingdom *Aza, a Hebrew roman ...
zoos, captives in North America are primarily ''O. tetrapis'' and the possibly unnamed West African species, but there are also some hybrids. Another study of individuals kept at
EAZA The European Association of Zoos and Aquaria (EAZA), is an organisation for the European zoo and aquarium community that links over 340 member organisations in 41 countries. EAZA membership is open to all zoos and aquaria across Europe that compl ...
zoos revealed a similar picture for Europe, but also that there were a few individuals of the fourth clade (native range in the wild unknown) and a single ''O. osborni''.


References


Further reading

*
''Osteolaemus tetraspis''@Crocodilian Species List
{{Taxonbar, from=Q244349 Crocodylidae Crocodilians of Africa Fauna of Central Africa Reptiles of West Africa dwarf crocodile Taxa named by Edward Drinker Cope